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Phylogeography of the Caribbean rock iguana (Cyclura): implications for conservation and insights on the biogeographic history of the West Indies.

The Caribbean rock iguana, Cyclura, has had an unstable intrageneric taxonomy and an unclear phylogenetic position within the family Iguanidae. We use mtDNA sequence data to address these issues and explore the phylogeographic history of the genus. ND4 to leucine tRNA sequence data were collected from multiple individuals of each of the eight species of Cyclura (including 15 of 16 subspecies) and from four localities of Iguana iguana (representative of this species' broad geographic range). This data set was combined with sequence data from Sites et al. (1996, Mol. Biol. Evol. 13, 1087-1105) and analyzed under maximum-parsimony and maximum-likelihood optimization criteria. The ND4 region provided good resolution for the majority of nodes, as indicated by high bootstrap support. In agreement with several recent molecular studies, Cyclura is recovered as monophyletic and is not closely related to any other genus, whereas Iguana is strongly supported as the sister taxon to Sauromalus. This result is statistically more likely than other published hypotheses of Iguanid relationships. Cyclura shows a southeast to northwest speciation sequence in the Caribbean, with the most ancient lineage on the Puerto Rican Bank. The amount of interspecific sequence divergence within Cyclura (maximum 11.4%) is very high in comparison to data from other iguanid taxa at this locus, suggesting that this group either has been in the Caribbean for a very long time or has gone through a very rapid rate of evolution at this locus. Using dates from other published studies, we calculate a molecular clock that suggests that Cyclura colonized the Caribbean between 15 and 35 mya. Several questions regarding subspecific taxonomy are raised in the analysis and await further investigation using a more rapidly evolving marker such as nuclear microsatellites.

Animals↗

Epistasis and the conversion of non-additive to additive genetic variance at population bottlenecks.

The effect of population bottlenecks on the mean and the additive variance generated by two neutral independent epistatic loci has been studied theoretically. Six epistatic models, used in the analysis of binary disease traits, were considered. Ancestral values in an infinitely large panmictic population were compared with their expectations at equilibrium, after t consecutive bottlenecks of equal size N (derived values). An increase in the additive variance after bottlenecks (inversely related to N and t) will occur only if the frequencies of the negative allele at each locus are: (1) low, invariably associated to strong inbreeding depression; (2) high, always accompanied by an enhancement of the mean with inbreeding. The latter is an undesirable property, making the pertinent models unsuitable for the genetic analysis of disease. For the epistatic models considered, it is unlikely that the rate of evolution may be accelerated after population bottlenecks, in spite of occasional increments of the derived additive variance over its ancestral value.

Animals↗

Transitions in understanding of RNA viruses: a historical perspective.

This chapter documents that RNA viruses have been known for over a century to be genetically variable. In recent decades, genetic and molecular analyses demonstrate that they form RNA quasispecies populations; the most rapidly mutating, highly variable and genetically versatile life forms on earth. Their enormous populations, rapid replication and extreme genetic plasticity can allow rates of evolution that exceed those of their eukaryotic host populations by millions-fold.

Evolution, Molecular↗

The adaptive significance of sexuality.

The theory predicts that inbred strains of mice are susceptible to tumor viruses that have escaped immune recognition. The theory implies routine evolutionary extinction of asexual species and that existing parthenogenetic species would have to have evolved recently from sexually reproducing precursors. This could be tested by determining the amount of DNA divergence between parthenogenetic and related sexually reproducing species. In mammals and birds genetic substitutions occur at comparable rates, but the AIDS virus evolves about 10 million times more rapidly. This confirms the basic imbalance of rates of evolution between microparasites and metazoan hosts, which is fundamental to the theory. The organization of most eukaryotic genes into exons and introns facilitates the generation of variety of gene products, and the molecular mechanism is similar to the mechanism that generates antibody diversity in the immune response and antigenic variation in trypanosomes. It has been proposed that noncoding 'selfish' DNA is the ultimate sexually transmitted disease. If this were the case, then asexually reproducing species would have an added advantage. An alternative hypothesis is proposed: noncoding DNA could provide insertion sites for retroviruses that would prevent them from being transcribed and replicated and thus moderate their proliferation much as absorber rods moderate proliferation of neutrons in a nuclear reactor. Flowering plants have pollen selection mechanisms that enforce heterozygosity at one or more loci. It has been proposed that analogous sperm selection mechanisms exist in mammals. Such a process would account for observation of a mysterious excess molecular divergence between different strains of inbred mice.

Adaptation, Physiological↗

RNA virus populations as quasispecies.

RNA virus mutation frequencies generally approach maximum tolerable levels, and create complex indeterminate quasispecies populations in infected hosts. This usually favors extreme rates of evolution, although periods of relative stasis or equilibrium, punctuated by rapid change may also occur (as for other life forms). Because complex quasispecies populations of RNA viruses arise probabilistically and differentially in every host, their compositions and exact roles in disease pathogenesis are indeterminate and their directions of evolution, and the nature and timing of "new" virus outbreaks are unpredictable.

Biological Evolution↗

The genetic consequences of primate social organization: a review of macaques, baboons and vervet monkeys.

Primates, as long-lived, iteroparous, socially complex mammals, offer the opportunity to assess the effects of behavior and demography on genetic structure. Because it is difficult to obtain tissue samples from wild primate populations, research in this area has largely been confined to terrestrial and semi-terrestrial old world monkeys (e.g., rhesus and Japanese macaques, vervets and several subspecies of baboons). However, these species display a multi-male, multi-female social structure commonly found in many other primate and non-primate mammals. Electrophoretic analyses of blood proteins from individually recognized and/or marked wild Himalayan rhesus monkeys, themselves the subject of long-term behavioral and demographic research, have begun to reveal the genetic consequences of such phenomena as social group fission, male-limited dispersion, non-consanguineous mating patterns, and agonistically defined male dominance. Specifically, rhesus social groups, consisting primarily of clusters of maternal relatives, appear to be non-random samples of a population's genotypes and genes. The genetic effects of social group fission are highly dependent on each group's size, demographic structure, and average degree of relatedness. In all cases fission contributes to the degree of intergroup genetic differentiation. Male-limited dispersion appears both to retard genetic differentiation between social groups and to lead to mating patterns that result in an avoidance of consanguinity. Groups, therefore, appear to be genetically outbred. Comparing these results with studies of other free-ranging or wild cercopithecines allows several generalizations: (a) genetic variation seems to be evenly distributed throughout each local population of multi-male social groups; (b) social groups, however, because they contain clusters of relatives, are distinctive in their specific frequencies of genes; (c) the degree of genetic differentiation between a population's social groups, because of the effects of social group fission and non-deterministic forms of male dispersal, is somewhat greater than expected on the basis of migration rates alone; and (d) the asymmetrical pattern of dispersion with respect to sex effectively precludes inbreeding in any one social group or the population as a whole. These observations have important implications for understanding the unusually rapid rates of evolution among the primates.

Animals↗

The Giardia lamblia actin gene and the phylogeny of eukaryotes.

The single-copy actin gene of Giardia lamblia lacks introns; it has an average of 58% amino acid identity with the actin of other species; and 49 of its amino acids can be aligned with the amino acids of a consensus sequence of heat shock protein 70. Analysis of the potential RNA secondary structure in the transcribed region of the G. lamblia actin gene and of the single-copy actin gene of nine other species did not reveal any conserved structures. The G. lamblia actin sequence was used to root the phylogenetic trees based on 65 actin protein sequences from 43 species. This tree is congruent with small-subunit rRNA trees in that it shows that oomycetes are not related to higher fungi; that kineto-platid protozoans, green plants, fungi and animals are monophyletic groups; and that the animal and fungal lineages share a more recent common ancestor than either does with the plant lineage. In contrast to small-subunit rRNA trees, this tree shows that slime molds diverged after the plant lineage. The slower rate of evolution of actin genes of slime molds relative to those of plants, fungi, and animals species might be responsible for this incongruent branching.

Actins↗

Interchromosomal gene conversion as a possible mechanism for explaining divergence patterns of ZFY-related genes.

The divergence pattern of mammalian ZFY-related genes from human (ZFY and ZFX) and mouse (Zfy-1 and Zfx) was reexamined on the basis of nucleotide substitutions at the synonymous codon-alternating positions. It is possible to explain the unusual divergence pattern of the mammalian Y-linked ZF genes by interchromosomal gene conversion by X-linked ZF genes. Furthermore, the rates of evolution of mammalian X- and Y-linked ZF genes were shown to agree well with those expected from our model.

Animals↗

Long term follow up of haemophilic arthropathy treated by Au-198 radiation synovectomy.

Intra-articular injection of Au-198 (gold synoviorthesis) has been used in the treatment of repeated haemarthroses of the elbows, knees, or ankles in 64 haemophiliacs. These patients had continued to have joint haemorrhages despite appropriate, episodic, substitutive therapy over a period of more than six months. Follow-up for an average period of 14 years in one knee in 38 male haemophiliacs showed 8 good, 23 fair, and 7 poor results with regard to their joint scores. It is concluded that Au-198 is an effective agent for radiation synovectomy, particularly in the early stages of the disease with minimal radiographic changes. It appears to reduce the incidence of haemarthrosis and to slow the rate of evolution of radiographic changes.

Adolescent↗

Quantitative subunit hybridization of drosophila alpha-glycerophosphate dehydrogenase.

The dimeric enzyme, alpha-Glycerophosphate dehydrogenase, was purified from eight Drosophila species by the method of Collier et al. (1976). The enzymes were inactivated at high pH and the conditions sufficient for reactivation were established. Electrophoretic patterns of reactivated alpha-glycerophosphate dehydrogenases which were mixed following inactivation of two species' enzymes, demonstrate that high pH dissociates the enzyme into its constituent subunits and reactivation involves subunit reassociation. Twenty interspecific combinations of dissociated enzymes were allowed to reassociate, and the amounts of both heterospecific and homospecific enzyme activity and protein were determined by densitometry. In all 20 tests there were no differences between observed and expected heterospecific:homospecific enzyme ratios. These results are consistent with the very slow rate of evolution of this enzyme in the family Drosophilidae (Collier and MacIntyre, 1977).

Animals↗

Evolutionary changes of alpha-crystallin and the phylogeny of mammalian orders.

The sequences of the A chains of the eye lens protein alpha-crystallin from seventeen mammalian species were compared. They showed a generally slow rate of evolution, but with marked variations in different lineages. Most substitutions have occurred in the C-terminal part of the chain, which probably forms part of the surface of the alpha-crystallin aggregate. The ancestral sequence method of Dayhoff revealed interesting indications about the phylogenetic relationships between the eleven mammalian orders that were represented by the investigated species. Most evident was the divergence of marsupial and placental orders. A notable resemblance between the hyrax and elephant sequences was observed, setting them apart from the ungulates, including whale. Primates, rodents, lagomorphs, insectivores and tupaiids seem to derive from a common stem group. These phylogenetic inferences are discussed in relation to current palaeontological and taxonomical opinions, and compared to evidence from other protein sequence data.

Amino Acid Sequence↗

Primitive haemoglobin.

The sequences of Petromyzon and Aplysia globins are compared with the postulated vertebrate and mollusc-vertebrate ancestors to see if differences exist in the rates of evolution of different types of residue positions. Between the mollusc-vertebrate ancestor and Aplysia globin there is no very striking pattern of changes except that the interior positions are relatively conserved. In the evolution of Petromyzon haemoglobin, the alpha 1 beta 2 contact area is relatively conserved. The homopolymeric binding of lamprey Hb seems to be a primitive function.

Animals↗

Inter- and intrapopulation studies of ancient humans.

For a genetic analysis of ancient human populations to be useful, it must be demonstrated that the DNA samples under investigation represent a single human population. Toward that end, we have analyzed human DNA from the Windover site (7000-8000 BP). MHC-I analysis, using allele-specific oligonucleotide hybridization to PCR amplified Windover DNA, microsatellite analysis by PCR of the APO-A2 repeat and mtD-loop 3' region sequencing on multiple individuals spanning nearly the full range of estimated burial dates all confirm the hypothesis that there is a persistence of both nuclear and mitochondrial haplotypes at Windover throughout its entire period of use. Thus, Windover can be considered a single population. Neighbor-joining tree analysis of mtDNA sequences suggests that some mitochondrial types are clearly related to extant Amerind types, whereas others, more distantly related, may reflect genetically distinct origins. A more complete sequence analysis will be required to firmly resolve this issue. Calibrating genetic relationships deduced by tree analysis, radiocarbon dates and burial position, yields a human mtD-loop DNA rate of evolution of 3700 to 14,000 years per percent change. Both values are within the range of recent, independently calculated values using estimates of evolutionary divergence or theoretical population genetics. Thus we are beginning to realize the promise of ancient DNA analysis to experimentally answer heretofore unapproachable questions regarding human prehistory and genetic change.

Alleles↗

Codon usage in muscle genes and liver genes.

Synonymous codon usage frequencies, derived from cDNA clone sequences, were compared for several sets of vertebrate genes. Gene sets as diverse as those expressed in avian skeletal muscle and in mammalian liver showed similar patterns of synonymous codon usage. There were no significant differences suggesting tissue-specific co-adaptation of codon usage patterns and tRNA anticodon profiles. The results indicate a consensus codon usage pattern for vertebrate genes which is largely independent of taxonomic class, tissue of expression, and the cellular fate and rate of evolution of the encoded proteins. Certain elements of the consensus codon usage pattern indicate that it is the product of natural selection and not simply a mutational equilibrium among phenotypically equivalent synonyms.

Animals↗

DNA hybridization evidence of hominoid phylogeny: results from an expanded data set.

The living hominoids are human, the two species of chimpanzees, gorilla, orangutan, and nine species of gibbons. The cercopithecoids (Old World monkeys) are the sister group of the hominoids. A consensus about the phylogeny of the hominoids has been reached for the branching order of the gibbons (earliest) and the orangutan (next earliest), but the branching order among gorilla, chimpanzees, and human remains in contention. In 1984 we presented DNA-DNA hybridization data, based on 183 DNA hybrids, that we interpreted as evidence that the branching order, from oldest to most recent, was gibbons, orangutan, gorilla, chimpanzees, and human. In the present paper we report on an expanded data set totaling 514 DNA hybrids, which supports the branching order given above. The ranges for the datings of divergence nodes are Old World monkeys, 25-34 million years (Myr) ago; gibbons, 16.4-23 Myr ago; orangutan, 12.2-17 Myr ago; gorilla, 7.7-11 Myr ago; chimpanzees-human, 5.5-7.7 Myr ago. The possible effects of differences in age at first breeding are discussed, and some speculations about average genomic rates of evolution are presented.

Animals↗

Evolutionary relationships of eukaryotic kingdoms.

The evolutionary relationships of four eukaryotic kingdoms--Animalia, Plantae, Fungi, and Protista--remain unclear. In particular, statistical support for the closeness of animals to fungi rather than to plants is lacking, and a preferred branching order of these and other eukaryotic lineages is still controversial even though molecular sequences from diverse eukaryotic taxa have been analyzed. We report a statistical analysis of 214 sequences of nuclear small-subunit ribosomal RNA (srRNA) gene undertaken to clarify these evolutionary relationships. We have considered the variability of substitution rates and the nonindependence of nucleotide substitution across sites in the srRNA gene in testing alternative hypotheses regarding the branching patterns of eukaryote phylogeny. We find that the rates of evolution among sites in the srRNA sequences vary substantially and are approximately gamma distributed with size and shape parameter equal to 0.76. Our results suggest that (1) the animals and true fungi are indeed closer to each other than to any other "crown" group in the eukaryote tree, (2) red algae are the closest relatives of animals, true fungi, and green plants, and (3) the heterokonts and alveolates probably evolved prior to the divergence of red algae and animal-fungus-green-plant lineages. Furthermore, our analyses indicate that the branching order of the eukaryotic lineages that diverged prior to the evolution of alveolates may be generally difficult to resolve with the srRNA sequence data.

Animals↗

Phylogenetic analysis of the isopenicillin-N-synthetase horizontal gene transfer.

A phylogenetic study of the isopenicillin-N-synthetase (IPNS) gene sequence from prokaryotic and lower eukaryotic producers of beta-lactam antibiotics by means of a maximum-likelihood approach has been carried out. After performing an extensive search, rather than invoking a global molecular clock, the results obtained are best explained by a model with three rates of evolution. Grouped in decreasing order, these correspond to A. nidulans and then to the rest of the eukaryotes and prokaryotes, respectively. The estimated branching date between prokaryotic and fungal IPNS sequences (852 +/- 106 MY) strongly supports the hypothesis that the IPNS gene was horizontally transferred from bacterial beta-lactam producers to filamentous fungi.

Aspergillus nidulans↗

A comparison of genetic variation in two sibling species pairs of haplodiploid insects.

Sibling species pairs of sweat bees (Halictus confusus and H. tumulorum) and pine sawflies (Neodiprion pratti and N. maurus) were surveyed for genetic variability using enzyme electrophoresis. Levels of heterozygosity were found to be within the ranges earlier recorded for Hymenoptera. Expected heterozygosities were not significantly higher in the sawflies than in the sweat bees. Estimates of genetic identity between the sibling species were not lower than those generally found for diplodiploid insect species: no evidence was found for an increased rate of evolution in these haplodiploids. Genetic identity data among populations of H. confusus and between Halictus species were within the range expected for conspecific populations and sibling species, respectively. In Neodiprion all genetic distances were low but the two populations of N. pratti had similar genetic distances as each did to N. maurus, indicating the necessity for further systematic studies of the genus.

Animals↗